The role of salal in forest regeneration problems in coastal British Columbia: problem or symptom?

The ericaceous shrub, salal (Gaultheria shallon Pursch) is associated with forest regeneration problems on a variety of site types along the west coast of North America. On dry sites, salal is a serious competitor for water; on wet (cedar-salal) sites, salal is thought to limit nutrient availability to trees by competing, interfering with mycorrhizae and short-circuiting the nitrogen cycle through production of phenolic compounds. Short-term field experiments and laboratory and greenhouse studies provided some evidence of each of these effects. However, long-term results from a suite of silvicultural trials and associated ecological studies indicated that salal is not the fundamental problem for regenerating conifers on cedar-salal sites. Fertilization of cedar-salal sites caused a large and sustained response in tree growth regardless of the presence of salal, and growth responses to salal removal were much smaller than fertilization responses. Greenhouse experiments indicated that salal does not have allelopathic effects on germination or growth of conifer seedlings. In laboratory studies, salal did not demonstrate a greater capacity to take up organic N forms than the conifers, and molecular studies uncovered a vast diversity of mycorrhizal fungi associated with salal and hemlock roots. Together these findings indicate that the nutrient short-circuiting hypothesis, based on assumptions about ericoid versus non-ericoid mycorrhizal plants does not adequately describe the nutrition of plants in these ecosystems. We conclude that salal should be viewed as a symptom of the underlying problem of poor nutrient supply on cedar-salal sites, and that fertilization, rather than salal control, is the optimal solution for improving forest regeneration on thesesites.